Large Herbivore Foraging and Ecological HierarchiesLandscape ecology can enhance traditional foraging theory

A pplications of optimal foraging theory (Pyke 1984, Schoener 1971) to large herbivores have been problematic (OwenSmith and Novellie 1981, Westoby 1974), partly because optimal foraging theory has been developed for predators and nectar feeders rather than large herbivores. Predators generally seek spatially scattered prey of nearly constant and high nutritional value. Large herbivores, on the other hand, confront an apparent food surplus, which is of low and highly variable nutritive quality (Belovsky 1984, Westoby 1978). Compared with the prey consumed by predatory animals, the food of large herbivores is much more likely to be widely dispersed over the landscape, rather than concentrated in discrete patches. Large herbivores interact with forage resources at several levels of ecological resolution. The animals confront a series of interrelated foraging problems, each on a different tempo-

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